Neotropical Ichthyology, 12(1):1-25, 2014 Copyright © 2014 Sociedade Brasileira de Ictiologia

Taxonomic revision of Nijssen & Isbrücker (: : ), with description of three new

Carine C. Chamon1,2 and Lúcia H. Rapp Py-Daniel3

A taxonomic review of Spectracanthicus Nijssen & Isbrücker, including Oligancistrus Rapp Py-Daniel, following a phylogenetic study, is presented. Additionally to S. punctatissimus (Steindachner) and S. murinus Nijssen & Isbrücker, three new species are recognized based on the examination of 159 specimens: S. immaculatus n. sp. from rio Tapajós basin, differs from its congeners by its color pattern consisting of a dark gray body, with no dots or spots, and by having very slender teeth; Spectracanthicus tocantinensis n. sp., from the rio Tocantins drainage is distinguished by the color pattern consisting of dark brown or black body with small, yellowish dots (except in S. punctatissimus), presence of thick teeth, infraorbital 4 forming most of the posterior edge of the orbit and the large basipterigium fenestrae; and Spectracanthicus zuanoni n. sp., from the rio Xingu basin is diagnosed by its color pattern consisting of large, white spots and by the larger orbital diameter. Other characters based on osteological features are also usefull to distinguish the species. A key to the species of the and a brief discussion of their threats and conservation are also provided. A revisão taxonômica de Spectracanthicus Nijssen & Isbrücker, incluindo Oligancistrus Rapp Py-Daniel, baseada em estudos filogenéticos recentes, é apresentada. Adicionalmente a S. punctatissimus (Steindachner) e S. murinus Nijssen & Isbrücker, três espécies novas são reconhecidas baseadas no exame de 159 espécimes: S. immaculatus, sp. n. proveniente da bacia do rio Tapajós, difere de suas congêneres pelo padrão de colorido do corpo cinza escuro sem pontos ou manchas e por possuir dentes bastante delgados; Spectracanthicus tocantinensis sp. n., da bacia do rio Tocantins pode ser distinguida pelo padrão de colorido do corpo marron escuro ou preto com pontos amarelados (exceto de S. punctatissimus), presença de dentes espessos, infraorbital 4 formando quase toda a margem posterior da órbita e pela presença de uma grande fenestra no basipterígio; e Spectracanthicus zuanoni sp. n., da bacia do rio Xingu é diagnosticada pelo padrão de colorido cinza escuro com grandes pontos brancos e por possuir maior diâmetro infraorbital. Outros caracteres osteológicos também podem ser utilizados para distinguir as espécies. Uma chave para as espécies do gênero e uma breve discussão sobre suas ameaças e conservação são também fornecidas. Key words: Acari fish, Aquarism, Amazon basin, Armored , Ornamental fishery.

Introduction divided into historically unstable subfamilies, most of them apparently monophyletic (Loricariinae, Hypoptopomatinae, The Loricariidae is the largest family within Siluriformes Lithogeninae, and Delturinae; Reis et al., 2006). However, the and includes more than 750 species, which represent about monophyly and composition of Hypostominae and Ancistrinae 25% of the diversity of (Reis et al., 2003; Ferraris, are still under scrutiny (Reis et al., 2003; Ferraris, 2007). 2007). The family is widely distributed in the Neotropical Different hypotheses of rearrangement of the Hypostominae region, from southeastern Costa Rica to northeastern Argentina have been proposed over the years (Regan, 1904; Gosline, (Isbrücker, 1980) and can be found in a large variety of 1947; Isbrücker, 1980; Schaefer, 1987; Armbruster, 2004), and freshwater environments. Loricariidae species are usually the subfamily was considered paraphyletic by several authors

1Museu de Zoologia da Universidade de São Paulo. Caixa Postal 04218-970 São Paulo, SP, Brazil. 2Universidade de Brasília - UnB, Câmpus Universitário Darcy Ribeiro, Instituto de Biologia - IB, Departamento de Zoologia, 70910-970 Brasília, DF, Brazil. [email protected] 3Instituto Nacional de Pesquisas da Amazônia, Coordenação de Pesquisas em Biologia Aquática., Manaus, AM, Brazil. [email protected] 1 2 Revision of Spectracanthicus

(Armbruster, 2004; Schaefer 1987; Montoya-Burgos et al., d’histoire naturelle de la Ville de Genève, Geneva (MHNG), 1998). The subfamily is currently recognized as comprising Museu Nacional, Universidade Federal do Rio de Janeiro, Rio the tribes Ancistrini (= Ancistrinae sensu Isbrücker, 1980), de Janeiro. (MNRJ), Museu de Zoologia da Universidade de São Hypostomini, Corymbophanini, Pterygoplichthyini, and Paulo, São Paulo (MZUSP), Naturhistorisches Museum Wien, (Armbruster, 2004). Although the Ancistrini is not Viena (NMW), Smithsonian Institution, National Museum of diagnosed by a unique characters (Armbruster, 2004), most Natural History, Washington D.C. (USNM), and Zoological of its genera are readily recognized by having the opercle Museum of Amsterdam (ZMA) now RMNH = Naturalis - modified into a rectangular bar- or sickle-shaped structure National Natuurhistorisch Museum, Leiden. with associated eversible plates supporting hypertrophied odontodes. However, this character is apparently reversed Spectracanthicus Nijssen & Isbrücker, 1987 in some taxa, and the opercle has triangular or oval shaped, which is the plesiomorphic state within Hypostominae (i.e., Spectracanthicus Nijssen & Isbrücker, 1987: 93.Type species: Spectracanthicus and ). Spectracanthicus murinus Nijssen & Isbrücker, 1987 by Spectracanthicus Nijssen & Isbrücker was established as original designation and monotypy. Gender masculine. a monotypic genus to include S. murinus from the rio Tapajós Oligancistrus Rapp Py-Daniel, 1989: 246. Type species: basin. Recently, the genus Oligancistrus Rapp Py-Daniel Chaetostomus punctatissimus Steindachner, 1881 by was proposed as a junior synonym of Spectracanthicus by original designation and monotypy. Gender masculine. Armbruster (2004) based in the argument of an unnecessary recognition of two monotypic closely related genera. Chamon Diagnosis. Spectracanthicus is diagnosed from all other (2012) further corroborates such hypothesis in a phylogenetic Ancistrini, except niveatus and B. beggini, study of the group, which resulted in Oligancistrus and by the presence of an expansion of as paraphyletic, given that Oligancistrus punctatissimus as dorsal-fin membrane connecting the last dorsal-fin ray of well as the new species fitting the diagnosis of that genus, to the adipose-fin spine vs.( dorsal fin not expanded and represented successive sister-groups of Spectracanthicus connected to adipose). Spectracanthicus is distinguished from murinus further justifying the synonymy proposed by Baryancistrus by having up to 25 teeth on premaxilla (vs. more Armbruster (2004). The genus is originally distinguished from than 25); and from Parancistrus by having small gill opening, 1 other Ancistrini by the presence of an expansion of the dorsal- reaching up to /3 of cleithrum length (vs. large gill opening, 1 fin membrane connecting it to the adipose-fin spine and the reaching nearly /2 of cleithrum length). presence of up to 25 teeth on premaxilla. Herein, a taxonomic revision of Spectracanthicus is presented. In addition, a brief Spectracanthicus murinus Nijssen & Isbrücker, 1987 phylogenetic discussion on the relationships and a key to the Figs. 1-2 species of the genus are provided. Spectracanthicus murinus Nijssen & Isbrücker, 1987: 94, fig. Material and Methods 1 [type locality: Poça de Pedra, rio Tapajós, Pará, Brazil]. - Burgess, 1989: 445 [citation]. - Eschmeyer, 1998: 1138 Measurements and counts were made according to Rapp Py- [catalog]. - Isbrücker, 2001: 32 [catalog]. - Isbrücker, Daniel & Zuanon (2005). Specimens were cleared and stained 2002: 29 [catalog]. - Fisch-Müller, 2003: 395 [catalog]. - (c&s) according to Taylor & Van Dyke (1985). Osteological Ferraris, 2007: 297 [catalog]. - Lujan et al., 2009: 50-56 nomenclature and vertebral counts follow Schaefer (1987) [citation]. - Camargo et al., 2012:169 [catalog, picture]. and Armbruster (2004). Vertebral counts include the first five vertebrae modified into Webberian apparatus, and the fused Diagnosis. Spectracanthicus murinus is distinguished from its pre-ural centrum is counted as a single element. Terminology congeners by the non-eversible cheek plate (vs. check plate for the laterosensory system of the head follows Arratia & eversible); the triangular-shaped opercle (vs. bar shaped); Huaquin (1995); homologies of the preopercle sensory canal presence of three branched anal-fin rays vs.( four); pale (yellow follow Schaefer (1988). Material examined was described in in life) bars on tip of dorsal and caudal fins vs.( bars absent; the following order: Institution number, count, measurements except in some specimens of S. punctatissimus); the frontal of minimum and maximum, locality, geographical coordinates, forming a small portion of orbit border (less than ¼ vs. frontal date of collection, and collectors. Some lots have missing data forming a large portion of orbit border); the anterior process concerning date of collection and/or collectors. Specimens of the pterotic-supracleithrum forming the posterior margin of examined belong to the following institutions: Instituto orbit (vs. pterotic-supracleithrum not forming posterior margin Nacional de Pesquisas da Amazônia, Manaus (INPA), Field of orbit in S. immaculatus or contacting only a small area of Museum of Natural History, Chicago (FMNH), Muséum orbit posterior margin in remaining species). Spectracanthicus C.C. Chamon & L.H.R. Py-Daniel 3

Fig. 1. Spectracanthicus murinus, holotype, MZUSP 22011, 59.7 mm SL, rio Tapajós, Pará State, Brazil. 4 Revision of Spectracanthicus

Fig. 2. Color in life of Spectracanthicus murinus (Image by L. M. Sousa). murinus can be further distinguished from S. immaculatus n. pterotic-supracleithrum with one median sized plate. sp. and S. zuanoni n. sp. by having pale (yellow in life) small Mouth moderate in size, nearly as long as wide. Lips large, dots on body and fins (vs. dots or spots absent, body and fins covered with papillae; size of papillae decreasing towards dark gray in S. immaculatus; or with large white spots in S. posterior margin of lower lip; central buccal papilla absent. zuanoni). It can be also distinguished from S. punctatissimus Upper lip folded over itself. Maxillary barbel short; base of and S. tocantinensis n. sp. by the lower counts of premaxillary barbel united to lips, with free tip. Lower lip not reaching and dentary teeth, 2 to 3 and 5 respectively (vs. 3 to 25 and 8 to anterior margin of coracoid. Medial end of premaxillary teeth 46 in S. punctatissimus; and 5 to 8 and 8-16 in S. tocantinensis). series curved inwards. Premaxillae (Fig. 3) and dentary narrow and elongate. Dentary strongly curved inwards. Teeth slightly Description. Morphometric and meristic data are summarized in thick, well developed, with long crown and large lateral cusp; Table 1. Dorsal profile of body slightly convex from tip of snout its distal edge slightly curved inward. Three to four pairs of to vertical through dorsal-fin origin; concave, nearly straight from well developed predorsal plates. Cheek plates not eversible; that point to caudal-fin origin. Ventral profile of body straight from without associated hypertrophied odontodes (Fig. 4). snout tip to caudal-fin origin. Ventral surface from tip of snout to Body covered by five longitudinal series of plates urogenital papillae lacking plates, except for few small plates at supporting odontodes. Keels absent. Three to four predorsal pectoral- and pelvic-fin origins. Greatest body width at pectoral plates; small azygous predorsal plates sometimes present girdle. Head and trunk lacking keels or ridges. Greatest body between predorsal plates. Eight furcate neural spines depth at dorsal-fin origin; lowest at caudal peduncle. supporting dorsal fin. Head wide, pointed anteriorlly; snout and cheek completely Dorsal-fin rays i,7, pterygiophores located posterior covered by numerous small plates, except for small naked to neural spines of vertebral centra 6-17. Dorsal-fin area on tip of snout. Snout slightly pointed in dorsal view. base very long, its length equivalent to 12 dorsal plates, Nasal L-shaped, elongate. Frontal short with a slight contact reaching pre-adipose plate; connected to adipose fin by with nares anteriorly and orbit posteriorly Anterior margin of thick membrane. Dorsal-fin spinelet V-shaped with lock frontal short, reaching posterior margin, or half length, of nare. mechanism. Pectoral and pelvic fins well developed, medial Parieto-supraoccipital short with posterior edge narrow, lacking portion much expanded relative to base; distal margin crest. Sphenotic short, contacting IO6; lacking conspicuous rounded. Pectoral-fin rays I,6; unbranched ray covered with odontodes. Orbit moderate in size (14.1-33.4% HL) placed conspicuous odontodes. Tip of adpressed pectoral fin almost dorsolaterally. Iris with small dorsal flap over pupil. Pterotic- reaching vertical through medial, unbranched, pelvic-fin supracleithrum short with few fenestrae; anterior process ray. Pelvic-fin rays i,5; pelvic-fin spine reaching vertical forming most posterior margin of orbit. Posterior area of through anal-fin base when adpressed. Anal-fin rays i,3, C.C. Chamon & L.H.R. Py-Daniel 5

Table 1. Morphometric and meristic data of Spectracanthicus murinus. Values are given as percents of standard length or head length. SD = standard deviation. Holotype n Min Max Mean SD MZUSP 22011 Standard length 59.7 28 44.1 96.7 - - Percents of Standard Length Head length 34.3 28 33.6 41.3 35.2 1.4 Head depth 18.1 28 16.6 22.5 19.2 1.7 Head width 31.3 28 26.8 31.9 29.2 1.2 Body depth 19.4 28 15.3 28.8 20.5 3.0 Body width at dorsal 28.3 28 25.0 31.1 27.8 1.7 Body width at anal 15.1 28 12.2 20.0 15.3 1.7 Predorsal length 43.5 28 40.8 45.9 43.7 1.1 Postdorsal length 24.1 28 22.0 63.7 51.6 15.7 Postanal length 30.2 28 28.5 38.6 32.3 2.3 Dorsal-spine length 26.2 28 0.0 31.7 26.9 5.5 Pectoral-spine length 29.7 28 27.5 31.6 29.6 1.1 Pelvic-spine length 27.8 28 25.0 30.0 27.7 1.4 Dorsal-base length 30.1 28 29.6 40.0 34.5 2.7 Caudal peduncle depth 16.4 28 11.6 16.4 13.4 1.2 Percents of Head Length Snout length 69.7 28 58.1 74.8 69.8 3.3 Interorbital width 34.7 28 31.4 39.5 36.0 1.9 Orbital diameter 15.1 28 14.1 33.4 18.2 3.4 Dentary length 9.3 28 4.3 9.6 8.0 1.4 Counts Mode Premaxillary teeth 3 28 2 3 3 - Dentary teeth 5 28 5 5 5 - Dorsal-fin rays i+7 28 i+7 i+7 i+7 - Pectoral-fin rays i+6 28 i+6 i+6 i+6 - Pelvic-fin rays i+5 28 i+5 i+5 i+5 - Anal-fin rays i+3 28 i+3 i+3 i+3 - Caudal-fin rays ii+14 28 ii+14 ii+14 ii+14 - Lateral line plates 21 28 21 22 22 - Caudal peduncle plates 8 28 7 8 7 - located posterior to haemal spines of vertebral centra 14- may present faint dots along body and fins. 17. Caudal fin i,14,i, truncate; supracaudal plates 7. Usually four procurrent caudal-fin rays. Caudal peduncle strongly Geographic distribution. Spectracanthicus murinus is known deep. Total vertebrae 26, precaudal 8-12. Sixth rib strongly from the rio Tapajós basin, near Itaituba and Santarém, Pará thickened, remaining ribs slender. Infraorbital with 7-8 State, Brazil (Fig. 5). pores. Infraorbital 4 with little contact with orbit posterior margin. Infraorbital 6 forming only postero-vental part of Fisheries and economical importance. Spectracanthicus orbit. Lateral line pores restricted to hypural plate. murinus is an important resource of ornamental fish. It is recognized by local fishermen and aquarists as “bicudo” Color in alcohol. Dorsal surface of body dark brown with (snouty). It is usually captured by diving with the aid of a numerous small, yellow dots regularly distributed along compressor, a typical way to capture ornamental fishes in head, body and fins. Most specimens with pale, bar-shaped Itaituba and Santarém regions (Sousa & Birindelli, 2009). The patch lacking chromatophores on caudal-fin distal tip. Ventral species citation to rio Xingu basin in Camargo et al., 2012 is surface ochre without dots. Long time preserved specimens probably a misidentification (H. Gimenes Junior, pers. comm.). 6 Revision of Spectracanthicus

Fig. 3. Palatine, maxillar, and premaxillar in (a) Spectracanthicus murinus, MZUSP 24293, 37.0 mm SL, (b) Spectracanthicus zuanoni, INPA 3957, 57.0 mm SL, (c) Spectracanthicus immaculatus, paratype, MZUSP 92617, 58.9 mm SL and (d) Spectracanthicus tocantinensis, paratype, MZUSP 34265, 65.4 mm SL. C.C. Chamon & L.H.R. Py-Daniel 7

Fig. 4. Suspensorium of Spectracanthicus murinus, MZUSP 24293, 37.0 mm SL. a. internal view; b. external view. Scale bars = 2 mm. Image by E. Baena. Material examined. Holotype. MZUSP 22011, 59.7, Brazil, Pará, São 1, 62.6 mm SL, Trairão, rio Jamaxim, ilha Terra Preta, 5°27’11”S Luis, rio Tapajós, 4°27’0.02”S 56°15’0.00”W, Expedição Permanente 55°52’40”W, 20 Oct 1991, L. Rapp Py-Daniel & J. Zuanon. INPA à Amazônia, H. A. Britski et al. Paratypes. All from Brazil, Pará, 26480, 1, 62.55 mm SL, Itaituba, obtained from ornamental fish rio Tapajós: MZUSP 21849, 2, 44.2-58.2 mm SL, São Luís, rio fishermen, 08 Nov 2006, L. M. Sousa & J. L. Birindelli. INPA 26483, Tapajós, Cachoeira Lombo de Anta, 4°27’60.00”S 56°14’60.00”W, 6, 44.4-84.8 mm SL, Itaituba, donate by fishmen of ornamental fish, Sep 1970, Expedição Permanente à Amazônia. MZUSP 27633, 1, 08 Nov 2006, L. M. Sousa & J. L. Birindelli. INPA 26488, 8, 60.4-96.7 45.0 mm SL, São Luis, rio Tapajós, cachoeira do Maranhãozinho, mm SL, Itaituba, above Itaituba and below cachoeiras de Pimental, 4°28’0.00”S 56°15’0.00”W, Expedição Permanente à Amazônia. 4°21’34”S 056°10’02”W, 08 Nov 2006, L. M. Sousa & J. L. Birindelli. ZMA 107.878, 2, 55.5-63.6 mm SL, São Luís, rio Tapajós, Cachoeira INPA 26507, 2, 4.7-57.6 mm SL, Pimental, corredeira do Pajaú e Lombo de Anta, 4°27’60”S 56°14’60”W, 6 Oct 1970, Expedição arredores, 4°35’04”S 56°15’32”W, 11 Nov 2006, L. M. Sousa & J. Permanente à Amazônia. ZMA 107.877, 1, 58.6 mm SL, São Luís, L. Birindelli. MZUSP 24293, 2, 36.1-39.1 mm SL (1 c&s, 36.1 mm poça de pedra no rio Tapajós, 8 Oct 1970. ZMA 107.876, 1, 44.1 SL) São Luís, cachoeira do Maranhãozinho, 4°28’S 56°15’W, 06-07 mm SL, Cachoeira do Maranhãozinho, rio Tapajós, near São Luis, Nov 1970, Expedição Permanente à Amazônia. MZUSP 34279, 13, 4°28’00”S 56°15’00”W, 06-07 Oct 1970, Expedição Permanente 19.0-46.2 mm SL (1 c&s, 36.8 mm SL) Pederneiras, below Itaituba, à Amazônia. Non-type. Brazil, Pará, rio Tapajós: INPA 6990, 4°16’41”S 55°59’09”W, 24 Out 1983, M. Goulding. 8 Revision of Spectracanthicus

Fig. 5. Distribution map for the species of Spectracanthicus. Circles, S. murinus; stars, S. punctatissimus; squares, S. immaculatus; triangle, S. tocantinensis and diamon, S. zuanoni. White symbols refers to type locality.

Spectracanthicus punctatissimus (Steindachner, 1881) Diagnosis. Spectracanthicus punctatissimus can be distinguished Figs. 6-7 from all congeners by having 12-14 vertebrae from first neural spine not including hypural plate (vs. 8-11 in remaining species). Chaetostomus punctatissimus Steindachner, 1881: 119. It further differs from its congeners (except from S. murinus) [Type locality: east of rio Xingu, Porto de Móz, Pará, by having yellowish bars on tip of dorsal and caudal fins; from Brazil, 1°45’S 52°10’W]. - Isbrücker & Nijssen, 1991: S. immaculatus and S. zuanoni by having small yellowish 349 [information about type locality, redescription]. - dots covering body (vs. color pattern uniform, with no dots or Eschmeyer, 1998: 1398 [catalog]. - Isbrücker, 2001: 26-30 large white spots, respectively); and from S. tocantinensis by [catalog]. having the rounded dorsal profile of snout vs.( snout pointed). Spectracanthicus punctatissimus. - Lujan et al., 2009:50-56 Spectracanthicus punctatissimus differs from S. murinus by [citation]. presence of a bar-shaped eversible opercle with conspicuous Parancistrus punctatissimus. - Eigenmann & Eigenmann, odontodes (vs. triangle-shaped opercle not eversible, without 1889: 44 [citation]. - Fowler, 1954: 172 [description]. - conspicuous odontodes) and by the presence of 4 unbranched Burgess, 1989: 435 [atlas]. - Isbrücker, 2002: 23 [catalog]. anal-fin rays vs( . 3 anal-fin unbranched rays). - Armbruster, 2004: 53 [phylogenetic analysis]. - Ferraris, 2007: 274 [catalog]. - Lujan et al., 2009: 50-56 [citation]. Description. Morphometric and meristic data summarized - Isbrücker, 1980: 58 [catalog]. in Table 2. Dorsal profile of body slightly convex from tip Hypostomus niveatus. - Eigenmann & Eigenmann, 1889: 44 of snout to vertical through dorsal-fin origin; concave nearly [citation]. straight from that point to dorsal-fin origin to caudal-fin origin. Oligancistrus punctatissimus. - Rapp Py-Daniel, 1989: 236 Ventral profile straight from snout tip to origin of caudal fin. [description of Oligancistrus]. - Fisch-Müller, 2003: 390 Ventral surface from tip of snout to urogenital papillae lacking [catalog]. - Ferraris, 2007:274 [catalog]. - Camargo et al., plates, except for few small plates at pectoral and pelvic-fins 2012:129 [catalog, picture]. origin. Greatest body width at pectoral girdle. Head and trunk C.C. Chamon & L.H.R. Py-Daniel 9

Fig. 6. Spectracanthicus punctatissimus, holotype, NMW 47206, 104.0 mm SL, rio Xingu, Pará State, Brazil. lacking keels or ridges. Greatest body depth at dorsal-fin lacking conspicuous odontodes. Orbit moderate in size (14.4- origin; lowest at caudal peduncle. 25.7% HL) placed dorsolaterally. Iris with small dorsal flap Head wide, convex dorsally; snout and cheeks completely over pupil. Pterotic-supracleithrum short with few fenestrae; covered by numerous small plates, except for small naked anterior process forming most posterior margin of orbit. area on tip of snout. Snout rounded in dorsal profile. Nasal Posterior area of pterotic-supracleithrum with one plate. elongated, L-shaped. Frontal short with a slight contact Mouth moderate in size, nearly as long as wide. Lips large, with nares anteriorly and orbit posteriorly. Anterior margin covered with papillae; size of papillae decreasing towards of frontal short, reaching posterior margin or half length of posterior margin of lower lip; central buccal papilla absent. nare. Parieto-supraoccipital short with posterior edge narrow, Upper lip folded over itself. Maxillary barbel short; base of lacking crest. Sphenotic short, without contact with IO6, barbel united to lips with free tip. Lower lip not reaching 10 Revision of Spectracanthicus

Fig. 7. Color in live of Spectracanthicus punctatissimus. Image by J. Birindelli. Table 2. Morphometric and meristic data of Spectracanthicus anterior margin of coracoid. Medial end of premaxillary teeth punctatissimus. Values are given as percents of standard length series curved inwards. Premaxillae and dentaries narrow and or head length. SD = standard deviation. elongate. Dentaries strongly curved inwards. Teeth medium Holotype n Min Max Mean SD in size, well developed, with long crown and large lateral NMW 47206 cusp, its distal edge slightly curved. Eversible cheek plates Standard length 104.0 53 33.7 119.8 - - with associated hypertrophied odontodes and disposed as Percents of Standard Length unique block connected to opercle, that can be everted to Head length 34.4 53 32.8 45.3 36.5 2.1 approximately 90o from head by opercle movements (Fig. 8). Head depth 19.4 53 15.9 27.9 20.4 2.5 Body covered by five longitudinal series of plates Head width 29.4 53 27.2 38.0 31.1 2.3 supporting odontodes. Keels absent. Three to four predorsal Body depth 17.8 53 15.1 29.8 20.9 3.3 plates; some small azygous predorsal plates sometimes present. Body width at dorsal 31.6 53 26.0 41.6 30.2 2.4 Eight neural bifid spines supporting dorsal fin. Dorsal-fin rays Body width at anal 14.9 53 12.1 21.1 15.9 1.9 i,7, located posterior to neural spines of vertebral centra 6-17. Predorsal length 42.6 53 36.4 49.1 43.7 2.1 Base of dorsal-fin base very long, its length equivalent to 12 Postdorsal length 31.0 53 18.7 67.5 57.3 12.9 dorsal plates, reaching pre-adipose plate and connected to Postanal length 32.3 53 24.9 35.8 30.6 2.4 adipose fin by thick membrane. Dorsal-fin spinelet V-shaped Dorsal-spine length 28.1 53 21.3 39.4 31.7 3.9 with lock mechanism. Pectoral and pelvic fins well developed, Pectoral-spine length 28.3 53 27.0 35.9 31.3 2.1 medial portion much expanded relative to base; distal margins Pelvic-spine length 28.0 53 22.9 32.4 28.7 2.1 rounded. Pectoral-fin rays I,6; unbranched ray covered with Dorsal-base length 28.8 53 30.2 49.3 37.9 3.4 conspicuous odontodes. Tip of adpressed pectoral fin almost Caudal peduncle depth 12.4 53 11.6 16.5 13.8 1.3 reaching vertical through medial, unbranched, pelvic-fin Percents of Head Length ray. Pelvic-fin rays i,5; pelvic-fin spine reaching vertical Snout length 66.0 53 53.9 76.1 62.6 4.7 through anal-fin base when adpressed. Anal-fin rays i,4 Interorbital width 37.1 53 27.4 40.4 33.4 2.8 located posterior to hemal spines of vertebral centra 14-17. Orbital diameter 17.1 53 14.4 25.7 20.3 2.7 Caudal-fin rays i,14,i, truncated; supracaudal plates. Five to six procurrent caudal-fin rays. Caudal peduncle strongly deep Dentary length 13.9 53 8.0 16.8 11.4 2.1 in lateral view. Total vertebrae 26, precaudal 8-12. Sixth rib Counts Mode strongly thickened, remaining ribs slender. Infraorbital with Premaxillary teeth 14 53 3 25 8 7-8 pores. Infraorbial 4 with little contact with orbit posterior Dentary teeth 25 53 8 46 25 - margin. Infraorbital 6 forming only postero-vental part of Dorsal-fin rays i+7 53 i+7 i+7 i+7 - orbit. Lateral line pores restricted to hypural plate. Pectoral-fin rays i+6 53 i+6 i+6 i+6 - Pelvic-fin rays i+5 53 i+5 i+5 i+5 - Color in alcohol. Dorsal surface of body dark brown to dark gray Anal-fin rays i+4 53 i+4 i+4 i+4 - with numerous, yellow, variable sized dots, from tiny to small, Caudal-fin rays ii+14 53 ii+14 ii+14 ii+14 - regularly distributed along head, trunk and fins. Density of dots Lateral line plates 22 53 20 22 22 - on body variable; pattern apparently independent of body size, Caudal peduncle 8 53 7 8 8 - however, juvenile specimens usually have fewer dots than adults. plates C.C. Chamon & L.H.R. Py-Daniel 11

Fig. 8. Suspensorium of Spectracanthicus punctatissimus, MZUSP 108570, 78.2 mm SL. (a) external view and (b) internal view. Scale bars = 2 mm. Image by E. Baena.

Pale bars on distal tips of dorsal and caudal fins usually present. Geographic distribution. The species is spread along the rio Irregular white stains rarely observed (called “mutants” specimens Xingu basin, near Altamira, Pará, Brazil. It occurs in sympatry by local fishermen). Ventral surface ochre, lacking dots. with Spectracanthicus zuanoni (Fig. 5).

12 Revision of Spectracanthicus

Fishery and economical importance. Spectracanthicus Coqueiro, Belo Monte, 03°06’54”S 051°43’15”W, 05 Nov 2008, punctatissimus is an important resource of ornamental fish. It L. Rapp Py-Daniel et al. MHNG 2546.100, 4, 32.84-53.23 mm SL, Balneário de Altamira, 3°12’32”S 52°12’17”W, 31 Jul-1 Aug is also recognized as L016 and L030 (L-number; Schraml & 1999, A. Werner. MHNG 2555.14, 16.4 -70.7 mm SL, Altamira, Schafer, 2004) by aquarists and ornamental fishermen. It is 3°12’32.74”S 52°12’17.58”W, Jul 1992, A. Werner. MZUSP usually captured by diving with aid of a compressor, a typical 107194, 3, 73.3-112.0 mm SL, bedrock, Jericoá, near Mucura way to capture ornamental fishes in Altamira, Pará. fall and the Mazinho farm, Altamira, 3°24’52”S 51°44’23”W, 07 Jun 2010, O. T. Oyakawa, J. Muriel-Cunha, C. C. Chamon, I. Fichberg, L. Rossi & A. Sawakushi (hereinafter Equipe ECIX Remarks. There is a great variation on premaxilla and dentary ). MZUSP 107206, 55, 35.1-86.1mm SL, Ilha de Babaquara, teeth counts (3-25 and 8-48, respectively), which seems to bedrock, Altamira, 3°24’11”S 52°12’27”W, 06 Jul 2010, Equipe be consequently related to a variability of the premaxilla and ECIX. MZUSP 107211, 6, 36.3-68.7 mm SL, Gorgulho da Rita, dentary length/width. Exhaustive examination of specimens Altamira, 3°20’26”S 52°11’4”W, 06 Jul 2010, Equipe ECIX. were unable to correlate that variation with sexual dimorphism, USNM 376496, 3, 72.1-82.7 mm SL, 72.5 km from Senador José Porfírio, 31.5 km from Porto do Moz, 2°01’59”S 52°14’54”W (no ontogeny, color pattern or geographical distribution along the information of date and collectors). river. Regardless of the variation of color and teeth counts, we prefer to sustain S. punctatissimus as single species, since these characteristics are overlapped within known populations. Spectracanthicus immaculatus, new species Figs. 9-10 Material examined. Holotype. NMW 47206, 103.96 mm SL, rio Xingu, Porto Moz, 1°44’55.10”S 52°14’17.04”W, 23 Ago-Sep Holotype. MZUSP 92797, 63.8 mm SL, Brazil, Pará, rio Tapajós, 1865, L. Agassiz & Mr. Vinhas. [Date of collection and collectors near Itaituba, approximately 4°16’46.56”S 55°59’5.77”W, 7 Nov information in Higuchi (1996)]. Non-types. All from Brazil, Pará, rio Xingu. INPA 3956, 8, (there is just one in the lot), 64.0 2006, L. M. Sousa & J. L. Birindelli. mm SL, cachoeira de Kaituká, Altamira, 3°12’0”S 52°12’0”W, 09 Oct 1990, L. Rapp Py-Daniel & J. Zuanon. INPA 3958 ,4, 20.9- Paratypes. MZUSP 92617, 11, 38.2-82.2 mm SL (1 c&s, 81.2 mm 54.2 mm SL, cachoeira de Kaituká, Altamira, 3°12’0”S 52°12’0”W, SL), Brazil, Pará, rio Tapajós near Itaituba and Pimental, approximately 09 Oct 1990, L. Rapp Py-Daniel & J. Zuanon. INPA 25786, 4°16’46.56”S 55°59’5.77”W, 8 Nov 2006, L. M. Sousa & J. L. Birindelli. 42, 3.6-42.7 mm SL, cachoeira do Kaituká, Altamira, 3°12’0”S 52°12’0”W, 09 Oct 1990, L. Rapp Py-Daniel & J. Zuanon. INPA 29304, 1, 70.9 mm SL, Costa do Junior, Altamira, Oct 1996, J. Diagnosis. Spectracanthicus immaculatus can be distinguished Zuanon. INPA 31416, 2, 33.7-110.8 mm SL, bedrock in front of from all congeners by its color pattern consisting of a dark gray the River, Comunidade do Maia, 3°30’44”S 51°44’43”W, 11 Oct body, without dots or spots, and by having very slender teeth 1996, J. Zuanon. INPA 31417, 3, 54.4-119.8 mm SL, bedrock in (vs. with small to mid-sized yellowish dots in S. punctatissimus, front of the River, Comunidade do Maia, 3°30’44”S 51°44’43”W, 09 Nov 2008, L. Rapp Py-Daniel et al. INPA 31424, 3, 87.4- S. murinus, and S. tocantinensis; and large white spots 110.9 mm SL, Cachoeira do Landi, 3°35’01”S 51°49’21”W, 08 in S. zuanoni). It further differs from Spectracanthicus Nov 2008, L. Rapp Py-Daniel et al. INPA 31446, 2, 46.3-57.4 punctatissimus and S. zuanoni by the pointed snout in dorsal mm SL, Comunidade do Maia, 3°31’42” S 52°45’02” W, 09 Nov view (vs. snout rounded). Other characters that differs the 2008, L. Rapp Py-Daniel et al. INPA 31453, 24, 34.9-79.2 mm new species from other congeners are: nasal quadrangular (vs. SL, near Altamira, Babaquara, 3°23’45”S 52°12’16”W, 07 Nov 2008, L. Rapp Py-Daniel et al. INPA 31461, 9, 36.5-69.7 mm nasal L-shaped); posterior margin of pterotic-supracleithrum SL, bedrock in front of the river, Itaobinha, Vitória do Xingu, without contact with posterior margin of orbit (vs. with large 3°23’45”S 52°12’16”W, 04 Nov 2008, L. Rapp Py-Daniel et al. contact with orbit in S. murinus and small contact in remaining INPA 31463, 2, 65.9-96.0 mm SL, Itaboinha, bedrock at the species); cartilage area of basipterygia short (vs. elongate in river, Vitória do Xingu, 2°53’22”S 51°56’26”W, 04 Nov 2008, L. remaining species, except in S. murinus); anterior fenestrae of Rapp Py-Daniel et al. INPA 31464, 3, 54.4-119.8 mm SL, ilha de Merencio (Pontão), Belo Monte, 3°06’17”S 51°43’33”W, 05 Nov cartilage area of basipterygia large in size (vs. small to median 2008, L. Rapp Py-Daniel et al. INPA 31486, 2, 79.5-114.8 mm size in remaining species, except in S. tocantinensis) (Fig. 11). SL, Arroz Cru, 3°25’20”S 51°57’22”W, 06 Nov 2008, L. Rapp Py- Daniel et al. INPA 31781, 5, 36.6-78.0, Coqueiro, Belo Monte, Description. Morphometric and meristic data summarized in 3°06’54”S 51°43’15”W, 05 Nov 2008, L. Rapp Py-Daniel et al. Table 3. Dorsal profile of body slightly convex from tip of snout INPA 31784, 9, 33.7-49.1 mm SL, Gorgulho da Rita, Babaquara, 3°20’14”S 52°11’18”W, 07 Nov 2008, L. Rapp Py-Daniel et al. to vertical through of dorsal fin; concave, nearly straight from that INPA 31785, 5, 47.0-119.0 mm SL, Paraná do Rio, Belo Monte, point to caudal-fin origin. Ventral profile straight from snout tip to 3°06’06”S 51°42’49”W, 05 Nov 2008, L. Rapp Py-Daniel et al. origin of caudal fin. Ventral surface from tip of snout to urogenital INPA 31782, 9, 59.4-62.3 mm SL, Itaboinha, bedrock at the papillae lacking plates, except for few small plates at pectoral and River, Vitória do Xingu, 3°53’22”S 51°56’26”W, 04 Nov 2008, pelvic-fins origins. Body deep and robust even at caudal peduncle. L. Rapp Py-Daniel et al. INPA 31783, 2, 41.3-77.4 mm SL, Gorgulho da Rita, Babaquara, 3°20’14”S 52°11’18”W, 07 Nov Head and trunk lacking keels or ridges. Greatest body depth at 2008, L. Rapp Py-Daniel et al. INPA 31786, 2, 42.2-50.0 mm SL, dorsal-fin origin; lowest at caudal peduncle. C.C. Chamon & L.H.R. Py-Daniel 13

Fig. 9. Spectracanthicus immaculatus, holotype, MZUSP 92797, 64.6 mm SL, rio Tapajós, Pará State, Brazil. 14 Revision of Spectracanthicus

Fig. 10. Color in live of Spectracanthicus immaculatus, paratype, MZUSP 92617, 81.2 mm SL. Image by L. Sousa.

Fig. 11. Basipterygia and pelvic fin of (a) Spectracanthicus murinus, MZUSP 24293, 37.0 mm SL; (b) S. immaculatus, paratype, MZUSP 92617; 58.9 mm SL and (c) S. tocantinensis, paratype, MZUSP 34265, 65.4 mm SL. Scale bars = 2 mm. Image by E. Baena. C.C. Chamon & L.H.R. Py-Daniel 15

Table 3. Morphometric and meristic data of Spectracanthicus IO6, lacking conspicuous odontodes. Orbit moderate in size immaculatus. Values are given as percents of standard length (13.6-16.2% HL), placed dorsolaterally in head. Iris with small or head length. SD = standard deviation. dorsal flap over pupil. Pterotic-supracleithrum short with few Holotype n Min Max Mean SD fenestrae; anterior process forming most posterior margin of MZUSP 92797 orbit. Posterior area of pterotic-supracleithrum with one plate. Standard length 64.6 11 38.2 82.2 69.0 - Mouth moderate in size, nearly as long as wide. Lips large, Percents of Standard Length covered with papillae; size of papillae decreasing towards Head length 23.5 11 34.1 36.2 35.4 0.7 posterior margin of lower lip; central buccal papilla absent. Head depth 11.5 11 16.6 29.4 20.4 3.1 Upper lip folded over itself. Maxillary barbel short; base of Head width 20.0 11 18.6 33.8 31.0 4.1 barbel united to lips with free tip. Lower lip not reaching anterior Body depth 11.2 11 16.6 23.1 20.1 1.9 margin of coracoid. Medial end of premaxillary teeth series Body width at 18.4 11 24.9 31.4 28.3 2.1 almost straight. Premaxillae and dentaries narrow and elongate. dorsal Dentaries strongly curved inwards. Teeth medium in size, well Body width at 8.7 11 12.5 16.8 14.0 1.2 developed, slender, with long crown and large lateral cusp. anal Distal edge of teeth slightly curved inward. Eversible cheek Predorsal length 27.1 11 39.8 44.3 42.1 1.0 plates with associated hypertrophied odontodes and disposed Postdorsal length 14.9 11 20.2 36.0 25.7 3.8 as unique block connected to opercle, that can be everted to Postanal length 21.0 11 24.4 32.8 28.2 2.0 approximately 90o from head by opercle movements (Fig. 12). Dorsal-spine 22.5 11 23.0 31.8 28.5 2.6 length Body covered by five longitudinal series of plates supporting odontodes. Keels absent. Three to four predorsal Pectoral-spine 21.0 11 27.8 35.1 29.3 1.9 length plates; some small azygous predorsal plates sometimes present. Pelvic-spine 19.8 11 24.4 27.8 26.1 0.9 Eight neural bifid spines supporting dorsal fin. Dorsal-fin rays length i,7, located posterior to neural spines of vertebral centra 7-17. Dorsal-base length 20.3 11 27.5 34.1 31.3 2.3 Dorsal-fin base very long, its length equivalent to 12 dorsal Caudal peduncle 10.2 11 12.7 14.3 13.5 0.5 plates, reaching pre-adipose plate and connected to adipose depth fin by thick membrane. Dorsal-fin spinelet V-shaped with Percents of Head Length locking mechanism. Pectoral and pelvic fins well developed, Snout length 14.4 11 55.2 63.7 60.0 2.8 medial portion much expanded relative to base; distal margins Interorbital width 7.5 11 30.0 38.2 33.4 2.1 rounded. Pectoral-fin rays I,6; unbranched ray covered with Orbital diameter 3.8 11 13.6 16.2 14.7 0.6 conspicuous odontodes. Tip of adpressed pectoral fin almost Dentary length 2.2 11 8.1 12.8 10.9 1.5 reaching vertical through medial, unbranched, pelvic-fin ray. Counts Mode Pelvic-fin rays i,5; pelvic-fin spine reaching vertical through Premaxillary teeth 5 11 6 22 12 - anal-fin base when adpressed. Anal-fin rays i,4 located Dentary teeth 7 11 8 30 18 - posterior to hemal spines of vertebral centra 14-17. Caudal-fin Dorsal-fin rays i+7 11 i+7 i+7 i+7 - rays i,14,i, truncated; supracaudal plates. Five to six procurrent Pectoral-fin rays i+6 11 i+6 i+6 i+6 - caudal-fin rays. Caudal peduncle strongly deep in lateral Pelvic-fin rays i+5 11 i+5 i+5 i+5 - view. Total vertebrae 26, precaudal 8-12. Sixth rib strongly Anal-fin rays i+4 11 i+4 i+4 i+4 - thickened, remaining ribs slender. Infraorbital with 7-8 pores. Caudal-fin rays ii+14 11 ii+14 ii+14 ii+14 - Infraorbital 4 with little contact with orbit by posterior margin. Lateral line plates 23 11 21 23 21 - Infraorbital 6 forming only postero-vental part of orbit. Lateral line pores restrict to hypural plate. Caudal peduncle 8 11 7 8 7 - plates Color in alchool. Ground color evenly dark gray to dark brown without dots or spots. Ventral surface ochre to light brown Head wide, convex dorsally; snout and cheeks completely without dots. Dorsal surface of body of live specimens gray. covered by numerous small plates, except for small naked area on tip of snout. Snout slightly rounded in dorsal profile. Distribution. Spectracanthicus immaculatus is known from rio Nasal elongated, L-shaped. Frontal short with a slight contact Tapajós basin, near Itaituba and Pimental, Pará State, Brazil. with nares anteriorly and orbit posteriorly. Anterior margin of frontal short, reaching posterior margin or half of length Etymology. The specific epithet immaculatus“ ” derives from of nare. Parieto-supraoccipital short with posterior edge Latin, meaning unspotted or unstained, in allusion to the coloration narrow, lacking crest. Sphenotic short, without contact with pattern of the species, lacking dots or spots. An adjective. 16 Revision of Spectracanthicus

Fig. 12. Suspensorium of Spectracanthicus immaculatus, (a) external view and (b) internal view, paratype, MZUSP 92617, 58.9 mm SL. Scale bars = 2 mm. Image by E. Baena.

Fishery and economical importance. Spectracanthicus slender in relation to its congeners. The number of premaxillary immaculatus is an important resource of ornamental fish. It is teeth (less than 25 teeth on each) is directly related to the bone also recognized by local fishermen and aquarists as “naná” or shape, which is narrow and elongated. Decrease of teeth counts “L363” (L-number; Schraml & Schafer, 2004; Werner et al. and narrowing of the premaxillare seems to be a derived condition 2005). Like S. murinus, it is also captured by diving with help within the Hypostominae, since the species usually have of a compressor, a typical way of capturing ornamental fishes larger and shorter premaxillae supporting more than 25 teeth. in Itaituba and Santarém regions (Sousa & Birindelli, 2009). Shape of teeth can vary greatly among the Hypostominae and even within Spectracanthicus. In some species of the Hypostomus Remarks. In Spectracanthicus immaculatus, teeth are extremely unicolor group, males in reproductive period acquire elongated, C.C. Chamon & L.H.R. Py-Daniel 17 slender and unicuspid teeth, which returns to its normal shape, rays (vs. three unbranched anal-fin rays). The new species shorter, thicker and with a small cuspid, after this period further differs from S. punctatissimus by the slightly pointed (Armbruster, 1997). However, this condition in S. immaculatus snout in dorsal view (vs. snout rounded in dorsal view). is not related to sexual dimorphism. On the other hand, in S. murinus the teeth are very short and thick, while the teeth have a Description. Morphometric and meristic data are summarized regular size and density in the remaining species. in Table 4. Dorsal profile of body slightly convex from tip of snout to vertical through of dorsal fin; concave, nearly straight Spectracanthicus tocantinensis, new species from that point to caudal-fin origin. Ventral profile straight Fig. 13 from snout tip to origin of caudal fin. Ventral surface from tip of snout to urogenital papillae lacking plates, except for few Holotype. MZUSP 110989, 49.5 mm SL; Brazil, Pará, Carreira small plates at pectoral and pelvic-fin origins. Body deep and Comprida, rio Itacaiunas, Tocantins drainage, 5°22’S 49° 07’W, M. robust even at caudal peduncle. Head and trunk lacking keels Goulding, 14 Oct 1983. or ridges. Greatest body depth at dorsal-fin origin; lowest at caudal peduncle. Paratypes. All from Brazil, Pará, rio Tocantins drainage. INPA 6045, Head wide, convex dorsally; snout and cheeks completely 7, 43.13-76.9 mm SL, poço d’água downstream of Tucuruí Dam, rio covered by numerous small plates, except for small naked Tocantins, 09 Oct 1984, Ichthyology staff, INPA (G. M. Santos & B. area on tip of snout. Snout slightly rounded in dorsal profile. Mérona). INPA 6049, 1, 61.8 mm SL, rio Tocantins, Jatobal, Tucuruí, Nasal elongated, L-shaped. Frontal short with a slight contact 05 Jul 1982, Ichthyology staff, INPA. INPA 6050, 1, 62.8 mm SL, rio with nares anteriorly and orbit posteriorly. Anterior margin Tocantins, Jatobal, Tucuruí, 17 Jul 2008, Ichthyology staff, INPA. INPA of frontal short, reaching posterior margin or half of length 2990, 140, 16.2-92.9 mm SL, rio Tocantins, downstream of Tucuruí of nare. Parieto-supraoccipital short with posterior edge Dam, 3°45’58”S 49°40’21”W, 31 Ago 1984, Ichthyology staff, INPA. narrow, lacking crest. Sphenotic short, without contact with INPA 11133, 2, 78.3-86.2 mm SL, rio Tocantins, Tucuruí, 01 Sep 1981, IO6, lacking conspicuous odontodes. Orbit moderate in size Ichthyology staff, INPA. FMNH 95555, 1, 64.0 mm SL, lagoon in (15.4-20.3% HL), placed dorsolaterally in head. Iris with small front of Jatobal, rio Tocantins, 4°34’00”S 49°39’00”W, 16 Sep 1970, dorsal flap over pupil. Pterotic-supracleithrum short with few Expedição Permanente à Amazônia. MNRJ 19347, 175, 36.0-85.3 mm fenestrae; anterior process forming most posterior margin of SL; rio Tocantins, near Tucuruí, Sep 1984, L. C. Alvarenga. MNRJ orbit. Posterior area of pterotic-supracleithrum with one plate. 19359, 29, 60.8-82.7 mm SL, rio Tocantins, near Tucuruí, Sep 1984, Mouth moderate in size, nearly as long as wide. Lips large, L. C. Alvarenga. MNRJ 19373, 7, 60.0-63.0 mm SL, rio Tocantins, covered with papillae; size of papillae decreasing towards near Tucurui, Sep 1984, L. C. Alvarenga. MZUSP 24125, 7, 56.4-74.0 posterior margin of lower lip; central buccal papilla absent. mm SL (1 c&s, 64.2 mm SL), Jatobal, lagoon in front of Jatobal, rio Upper lip folded over itself. Maxillary barbel short; base of Tocantins, 49°39’00”W 4°34’00”S, 16 Sep 1970, Expedição permanente barbel united to lips with free tip. Lower lip not reaching à Amazônia. MZUSP 34265, 11, 38.7-72.0 mm SL (1 c&s, 65.4 mm anterior margin of coracoid. Medial end of premaxillary teeth SL), same locality as holotype. MZUSP 39927, 1, 64.5 mm SL, lagoon series almost straight. Premaxillae and dentaries narrow and in front of Jatobal, rio Tocantins, 4°34’00”S 49°39’00”W, 16 Sep 1970, elongate. Dentaries strongly curved inwards. Teeth medium Expedição Permanente à Amazônia. in size, well developed, slender, with long crown and large lateral cusp. Distal edge of teeth slightly curved inward Diagnosis. Spectracanthicus tocantinensis can be diagnosed inward. Eversible cheek plates with associated hypertrophied from its congeners by the infraorbital 4 forming most of the odontodes and disposed as unique block connected to opercle, posterior edge of the orbit (vs. IO4 forming a minute portion that can be everted to approximately 90o from head by opercle of orbital rim) and by the large basipterigium fenestrae (vs. movements (Fig. 12). medium or small-sized basipterigium fenestrae) (Fig.11). It can Body covered by five longitudinal series of plates be further distinguished from S. immaculatus and S. zuanoni supporting odontodes. Keels absent. Three to four predorsal by its color pattern, consisting of small, yellowish dots, and by plates; some small azygous predorsal plates sometimes present. the presence of thick teeth (vs. dots lacking in S. immaculatus Eight neural bifid spines supporting dorsal fin. Dorsal-fin rays and large, white spots in S. zuanoni; and slender teeth). It i,7, located posterior to neural spines of vertebral centra 6-17. is additionally distinguished from S. zuanoni by the lower Dorsal-fin base very long, its length equivalent to 12 dorsal number of dentary teeth (8-16 vs.19-43). Spectracanthicus plates, reaching pre-adipose plate and connected to adipose fin tocantinensis differs from S. murinus by the presence of a by thick membrane. Dorsal-fin spinelet V-shaped with locking bar-shaped, eversible opercle with conspicuous odontodes mechanism. Pectoral and pelvic fins well developed, medial (vs. triangle shaped opercle not eversible, without conspicuous portion much expanded relative to its base; distal margins odontodes) and by the presence of four unbranched anal-fin rounded. Pectoral-fin rays I,6; unbranched ray covered with 18 Revision of Spectracanthicus conspicuous odontodes. Tip of adpressed pectoral fin almost Color in alcohol. Dorsal surface of body and fins dark brown reaching vertical through medial, unbranched, pelvic-fin ray. with small to median-sized yellowish dots regularly distributed Pelvic-fin rays i,5; pelvic-fin spine reaching vertical through along head, trunk and fins. Body dots sparced and usually anal-fin base when adpressed. Anal-fin rays i,4 located fewer in juveniles. Ventral surface light tan without dots. posterior to hemal spines of vertebral centra 14-17. Caudal- fin rays i,14,i, truncated; seven supracaudal plates. Five to six Geographic distribution. Spectracanthicus tocantinensis is procurrent caudal-fin rays. Caudal peduncle strongly deep in known from the lower rio Tocantins, near Tucuruí and Jatobal, lateral view. Total vertebrae 26, precaudal 8-12. Ribs slender, Pará, Brazil (Fig. 5). except strong sixth rib. Infraorbital with 7-8 pores. Infraorbital 4 with great contact with orbit by posterior margin. Infraorbital Etymology. The specific epithet tocantinensis“ ” is in allusion 6 formed just postero-vental part of orbit. Lateral line pores to the type locality of the new species, the rio Tocantins. An restrict to hypural plate. adjective.

Fig. 13. Spectracanthicus tocantinensis, holotype, MZUSP 110989, 56.4 mm SL, rio Tocantins, Pará State, Brazil. Image by E. Baena. C.C. Chamon & L.H.R. Py-Daniel 19

Table 4. Morphometric and meristic data of Spectracanthicus 51°53’0”W, 05 Oct 1990, J. Zuanon. INPA 25874, 3, 69.9-126.2 mm tocantinensis. Values are given as percents of standard length SL, ilha do Sr. Izaltino, bedrock beside the island (left margim), or head length. SD = standard deviation. 1°30’0”S 51°53’0”W, 14 Sep 1997, J. Zuanon. INPA 25876, 1, 123.1 Holotype n Min Max Mean SD mm SL, 1°30’0”S 51°53’0”W. INPA 25877, 2, 70.6-129.3 mm SL, MZUSP 110989 ilha do Sr. Izaltino, left margin, 1°30’0”S 51°53’0”W, 14 Sep 1997, J. Zuanon. INPA 25878, 1, 107.3 mm SL, 1°30’0”S 51°53’0”W. INPA Standard length 56.4 31 57.7 93.3 72.5 8.7- 25879, 2, 76.9-112.8 mm SL, ilha do Sr. Izaltino, bedrock beside the Percents of Standard Length island (left margim), 1°30’0”S 51°53’0”W, 14 Sep 1997, J. Zuanon. Head length 20.7 31 32.4 37.3 35.4 1.4 INPA 25881, 1, 93.2 mm SL, ilha do Bananal, Senador José Porfírio, Head depth 11.2 31 17.4 28.1 20.5 2.2 08 Sep 1997, J. Zuanon. INPA 25882, 2, 55.9-82.8 mm SL, corredeiras Head width 18.6 31 30.2 34.8 32.0 1.2 do Arini, 1°30’0”S, 51°53’0”W, 07 Sep 1997, J. Zuanon. MZUSP Body depth 11.2 31 17.3 26.4 20.6 2.3 107200, 3, 59.4-86.1 mm SL, bedrock near Jericoá, near Mucura fall Body width at dorsal 15.6 31 25.9 31.9 28.1 1.5 and above Mazinho’s farm, Altamira, 3°24’52”S 51°44’23”W, 07 Jun Body width at anal 8.3 31 14.6 19.1 16.1 1.3 2010, Equipe ECIX (O. T. Oyakawa, J. Muriel-Cunha, C. C. Chamon, Predorsal length 24.9 31 42.3 48.8 43.9 1.5 I. Fichberg, L. Rossi & A. Sawakushi). MZUSP 107202, 3, 37.8-95.3 Postdorsal length 13.5 31 16.3 28.0 22.1 2.9 mm SL, Gorgulho da Rita, Altamira, 3°20’26”S 52°11’4”W, 06 Jul Postanal length 15.5 31 25.1 35.4 29.6 3.2 2010, Equipe ECIX. MZUSP 107207, 17, 41.3-98.1 mm SL, ilha Dorsal-spine length 15.7 31 25.5 31.8 28.7 1.7 de Babaquara, bedrock, Altamira, 3°24’10”S 52°12’27”W, 06 Jul Pectoral-spine length 16.6 31 27.3 31.9 29.4 1.2 2010, Equipe ECIX. Pelvic-spine length 15.8 31 24.2 28.7 26.8 1.2 Dorsal-base length 17.5 31 30.6 36.2 33.3 1.7 Non-Types. All from Brazil, Pará, rio Xingu. INPA 3957, 5, Caudal peduncle depth 8.2 31 14.0 18.0 14.9 1.0 20.1-58.8 mm SL, cachoeira de Baituká, 1°30’0”S 51°53’0”W, 09 Percents of Head Length Oct 1990, L. Rapp Py-Daniel & J. Zuanon. INPA 3966, 2, 67.1- Snout length 12.9 31 59.3 70.8 63.5 2.6 96.6 mm SL, Praia do Pedral, 1°30’0”S 51°53’0”W 07 Oct 1990, Interorbital width 7.0 31 31.7 37.7 34.8 1.4 L. Rapp Py-Daniel & J. Zuanon. INPA 28545, 2, 27.5-23.8 mm Orbital diameter 3.6 31 15.4 20.3 17.9 1.2 SL, Costa do Junior, 03°29’28”S 52°19’07”W, 23 Sep 1997, J. A. Dentary length 1.8 31 8.6 15.8 11.5 2.1 Zuanon. INPA 31411, 4, 78.4-92.9 mm SL, bedrock in front of the Counts Mode River, Comunidade do Maia, 03°30’44”S 51°44’43”W, 09 Nov 2008, L. Rapp Py-Daniel et al. INPA 31422, 3, 59.3-111.3 mm SL, Premaxillary teeth 7 31 5 8 6 - cachoeira do Landi, 03°35’01”S 51°49’21”W, 08 Nov 2008, L. Dentary teeth 10 31 8 16 12 - Rapp Py-Daniel et al. INPA 31454, 2, 47.6-71.1 mm SL, Gorgulho Dorsal-fin rays i+7 31 i+7 i+7 i+7 - da Rita, 03°20’14”S 52°11’18”W, 07 Nov 2008, L. Rapp Py-Daniel Pectoral-fin rays i+6 31 i+6 i+6 i+6 - et al. INPA 31455, 6, 41.4-72.4 mm SL, Babaquara, 03°23’45”S Pelvic-fin rays i+5 31 i+5 i+5 i+5 - 52°12’16”W, 07 Nov 2008, L. Rapp Py-Daniel et al. INPA 31460, 8, Anal-fin rays i+4 31 i+4 i+4 i+4 - 35.3-89.8 mm SL, Itaboinha, bedrock, Vitória do Xingu, 03°53’22”S Caudal-fin rays ii+14 31 ii+14 ii+14 ii+14 - 51°56’26”W, 04 Nov 2008, L. Rapp Py-Daniel et al. INPA 31480, Lateral line plates 23 31 20 24 23 - 11, 53.1-126.8 mm SL, Arroz Cru, 03°25’20”S 51°57’22”W, 06 Nov Caudal peduncle plates 7 31 7 8 7 - 2008, L. Rapp Py-Daniel et al. INPA 31775, 1, 113.0 mm SL, ilha do Davi, Altamira, 03°32’39”S 51°57’29”W, 01 Nov 2008, L. Rapp Spectracanthicus zuanoni, new species Py-Daniel et al. INPA 31794, 2, 50.5-54.0 mm SL, Belo Monte, Figs. 14-15 03°06’54”S 51°43’15”W, 05 Nov 2008, L. Rapp Py-Daniel et al. MHNG 2578.027, 1, 81.8 mm SL, Altamira (?), imported from Belém, Oligancistrus sp. - Werner et al., 2005: 23 [catalog; picture]. D. Fisher. MHNG 2.680.014, 1, 79.9 mm SL, aquarium import, R. Oligancistrus n. sp. 2. - Camargo et al., 2012: 131 [catalog; Covain. MHNG 2684.026, 2, 62.2-66.6 mm SL, aquarium import, picture]. R. Covain.

Holotype. INPA 25874, 122.4 mm SL, ilha do Sr. Izaltino, rio Xingu, Diagnosis. Spectracanthicus zuanoni can be diagnosed Altamira, Pará, Brazil, 3°16’21”S 52°12’7”W, J. Zuanon, 14 Sep 1997. from all other congeners by its color pattern consisting of large, white spots (vs. small yellowish dots in S. murinus, S. Paratypes. All from Brazil, Pará, rio Xingu. INPA 4031, 2, 78.2- punctatissimus, and S. tocantinensis, and dark gray lacking 127.9 mm SL, 1°30’0”S 51°53’0”W, 05 Oct 1990, L. Rapp Py-Daniel dots in S. immaculatus). It also differs from congeners, except & J. Zuanon. INPA 4147, 1, 84.9 mm SL, ilha de Babaquara, 1°30’0”S S. murinus, by the larger orbital diameter (up to 29.2% of HL 20 Revision of Spectracanthicus

Fig. 14. Spectracanthicus zuanoni, holotype, INPA 25874, 122.5 mm SL, rio Xingu, Pará State, Brazil. vs. up to 25.7% in S. punctatissimus, 20.3% in S. tocantinensis, by present high counts of dentary teeth (19-43 vs. 8-16) and 16.2% in S. immaculatus). It further differs from S. murinus by by IO4 forming just a small area of orbit (vs. IO4 forming the rounded dorsal view of the snout, the bar-shaped, eversible posterior edge of orbit almost totally). opercle with conspicuous odontodes, and the presence of four unbranched anal-fin rays vs.( snout slightly pointed; opercle Description. Morphometric and meristic data summarized triangle-shaped, not eversible, lacking conspicuous odontodes; in Table 5. Dorsal profile of body slightly convex from tip of three anal-fin unbranched rays). FromS. tocantinensis it differs snout to vertical through of dorsal fin; concave, nearly straight C.C. Chamon & L.H.R. Py-Daniel 21

Fig. 15. Color in live of Spectracanthicus zuanoni. Image by J. Birindelli.

from that point to caudal-fin origin. Ventral profile straight everted to approximately 90o from head by opercle movements. from snout tip to origin of caudal fin. Ventral surface from tip Body covered by five longitudinal series of plates of snout to urogenital papillae lacking plates, except for few supporting odontodes. Keels absent. Three to four predorsal small plates at pectoral and pelvic-fins origins. Body deep and plates; some small azygous predorsal plates sometimes robust even at caudal peduncle. Head and trunk lacking keels present. Eight neural bifid spines supporting dorsal fin. Dorsal- or ridges. Greatest body depth at dorsal-fin origin; lowest at fin rays i,7, located posterior to neural spines of vertebral caudal peduncle. centra 6-17. Dorsal-fin base very long, its length equivalent to Head wide, convex dorsally; snout and cheeks completely 12 dorsal plates, reaching pre-adipose plate and connected to covered by numerous small plates, except for small naked adipose fin by thick membrane. Dorsal-fin spinelet V-shaped area on tip of snout. Snout slightly rounded in dorsal profile. with lock mechanism. Pectoral and pelvic fins well developed, Nasal elongated, L-shaped. Frontal short with a slight contact medial portion much expanded relative to base; distal margins with nares anteriorly and orbit posteriorly. Anterior margin of rounded. Pectoral-fin rays i,6; unbranched ray covered with frontal short, reaching posterior margin or half of length of conspicuous odontodes. Tip of adpressed pectoral fin almost nare. Parieto-supraoccipital short with posterior edge narrow, reaching vertical through medial, unbranched, pelvic-fin ray. lacking crest. Sphenotic short, without contact with IO6, Pelvic-fin rays i,5; pelvic-fin spine reaching vertical through lacking conspicuous odontodes. Orbit enlarged (16.6-29.2% anal-fin base when adpressed. Anal-fin rays i,4 located HL), placed dorsolaterally in head. Iris with small dorsal flap posterior to hemal spines of vertebral centra 15-17. Caudal-fin over pupil. Pterotic-supracleithrum short with few fenestrae; rays i,14,i, truncated; supracaudal plates. Five to six procurrent anterior process forming most posterior margin of orbit. caudal-fin rays. Caudal peduncle strongly deep in lateral Posterior area of pterotic-supracleithrum with one plate. view. Total vertebrae 26, precaudal 8-12. Sixth rib strongly Mouth moderate in size, nearly as long as wide. Lips large, thickened, remaining ribs slender. Infraorbital with 7-8 pores. covered with papillae; size of papillae decreasing towards Infraorbital 4 with little contact with orbit by posterior margin. posterior margin of lower lip; central buccal papilla absent. Infraorbital 6 formed just postero-vental part of orbit. Lateral Upper lip folded over itself. Maxillary barbel short; base of line pores restrict to hypural plate. barbel united to lips with free tip. Lower lip not reaching anterior margin of coracoid. Medial end of premaxillary teeth Color in alcohol. Dorsal surface of body and fins evenly series almost straight. Premaxillae and dentaries narrow and brown covered by large white spots. Spots more frequent elongate. Dentaries strongly curved inwards. Teeth medium and conspicuous in dorsal and caudal fins than in pectoral, size, well developed, slender, with long crown and large lateral pelvic and anal fins. Some specimens, especially juveniles, cusp. Distal edge of teeth slightly curved inward. Eversible can present overlapped spots forming smudges. Head poorly cheek plates with associated hypertrophied odontodes and spotted, some individuals can present vermicular bands near disposed as unique block connected to opercle, that can be snout region. Abdomen region ochre without spots. 22 Revision of Spectracanthicus

Table 5. Morphometric and meristic data of Spectracanthicus L354 (L-number; Schraml & Schafer, 2004) by aquarists and zuanoni. Values are given as percents of standard length or local fishermen. As in Itaituba and Santarém, at rio Tapajós, head length. SD = standard deviation. Altamira is also known for its ornamental fishery activities.

Holotype n Min Max Mean SD Ecological notes. Spectracanthicus zuanoni is a nocturnal INPA 25874 species that feeds on algae and other food items off periphyton Standard length 122.5 32 59.3 129.0 92.5 - by grazing over rock bottom. According to Zuanon (1999) and Percents of Standard Length Rapp Py-Daniel & Zuanon (2005), Spectracanthicus zuanoni Head length 44.4 32 32.7 42.1 36.0 1.9 is a moderate reophilic species that is found in rock-bottom Head depth 26.2 32 18.1 29.1 21.7 2.6 areas subjected to backwater and strong currents. Specimens Head width 41.1 32 28.1 38.2 32.5 2.1 were found individually or in groups of three, under shelter Body depth 27.9 32 19.5 28.5 23.0 2.7 boulders (adults) and spaces beneath rocks (juveniles), Body width at dorsal 37.3 32 27.4 38.1 31.3 2.3 up to 2m deep. The juveniles share the shelters with other Body width at anal 21.5 32 12.7 22.7 16.8 2.2 loricariids such as sp., A. ranunculus, Baryancistrus Predorsal length 54.2 32 38.7 49.4 43.4 2.3 xanthellus, Spectracanthicus punctatissimus, Postdorsal length 26.4 32 19.2 27.7 22.7 2.0 tricornis, , vittata, and Postanal length 36.7 32 29.9 35.2 32.2 1.4 aff. barbatus. Dorsal-spine length 39.3 32 28.0 36.7 32.5 3.0 Pectoral-spine length 43.8 32 29.2 35.9 32.6 1.9 Discussion Pelvic-spine length 36.4 32 23.6 33.6 29.1 2.5 Dorsal-base length 44.4 32 20.5 41.7 34.2 4.4 Isbrücker (1980) and Schaefer (1986, 1987) diagnosed Caudal peduncle 19.7 32 14.5 20.8 16.8 1.7 the Ancistrini (sensu Armbruster, 2004) according to depth the presence of evertible cheek plate, usually bearing Percents of Head Length hypertrophied odontodes, a feature also present in Snout length 30.0 32 58.9 70.4 64.1 3.2 Pterygoplichthyini. However, according to Armbruster (2004), Interorbital width 13.6 32 44.6 62.2 52.9 4.6 the Pterygoplichthyini are distinguished from Ancistrini the Orbital diameter 7.7 32 16.6 29.2 23.5 3.5 by the presence of a connective tissue covering the stomach. Dentary length 6.2 32 15.0 29.0 20.8 3.9 Characters like the eversible cheek plates and presence of four Counts Mode umbranched anal-fin rays, present in most of Ancistrini taxa, Premaxillary teeth 14 32 6 25 14 - are both absent in Spectracanthicus murinus, due to reversion. Dentary teeth 30 32 19 43 30 - The most evident character that distinguishes Dorsal-fin rays i+7 32 i+7 i+7 i+7 - Spectracanthicus from other loricariids is the membranous connection between dorsal and adipose fins. Although such Pectoral-fin rays i+6 32 i+6 i+6 i+6 - feature is present in other loricariid genera (i.e., Baryancistrus Pelvic-fin rays i+5 32 i+5 i+5 i+5 - and Parancistrus), other characters readily distinguish these Anal-fin rays i+4 32 i+4 i+4 i+4 - from Spectracanthicus: the mesial wall of the metapterygoid Caudal-fin rays ii+14 32 ii+14 ii+14 ii+14 - being much taller than the lateral; the spoon-shaped anterior Lateral line plates 23 32 21 25 22 - process of the metapterygoid; and the presence of a deep Caudal peduncle 8 32 7 8 7 - plates pouch on the lateral ethmoid (Armbruster, 2004). Chamon (2012) proposed four additional synapomorphies supporting the monophyly of the genus: the presence of a cartilaginous Geographic distribution. Spectracanthicus zuanoni is known second basibranchial; the pointed distal margin of the Weberian from the rio Xingu basin, near Altamira and Belo Monte, Pará apparatus transverse process; the short hypurapophysis; and a State, Brazil (Fig. 5). great hiatus between the inferior hypural lobe and last hemal spine. All synapomorphies proposed in the aforementioned Etymology. The specific epithet is in honor to Jansen Zuanon studies were observed in all species of Spectracanthicus (INPA) due to his contribution to the knowledge of Neotropical recognized herein. In addition to these features, all species Ichthyology. Jansen Zuanon was also the first ichthyologist of Spectracanthicus (except S. immaculatus) share the who collected and identified this species as a new one. contact between sphenotic and IO6; the elongate anterior metapterygoid process; and the enlogate lateropterygium, Fishery and economical importance. Spectracanthicus not surpassing the vertical through the anterolateral process zuanoni is also recognized as “acari bola branca”, L020 and of the basipterygia. C.C. Chamon & L.H.R. Py-Daniel 23

Although species of Spectracanthicus can be distinguished direct decrease of stocks by fishing. Among fish species that from each other based on morphological features, they are endangered, only a small number of these have commercial can be promptly diagnosed from each other by their color interest as ornamental. In relation to rio Xingu ichthyofauna, pattern. In fact, the color pattern is a very useful source of it is forbidden to capture some species for ornamental purpose characters allowing the identification of loricariid fishes, when the decline of these populations is remarkable, for but some intraspecific variation is often common, (as with instance zebra that was considered endangered other characters sources) and must be considered. Such according to Normative Instruction MMA 05/2004 of the plasticity has rendered fishermen and aquarists to recognize Brazilian Goverment. specimens, falling into that variation, as different species. For the majority of the ornamental loricariid fish species Specimens of Spectracanthicus puntatissimus, for instance, from Amazon basin, the most imminent danger is the are recognized under different L numbers (Schraml & construction of dams. In the case of rio Xingu, the project Schaefer, 2004) according to the amount and size of the spots, of Belo Monte Dam includes a major barrage located 40 km and its occurrence along the rio Xingu: L016 (specimens from downstream from the city of Altamira (Volta Grande region), Altamira); L030 (specimens from Belo Monte); and L353 forming the reservoir Xingu. As a consequence, areas of (specimens from the rio Iriri). Some other characters referring the River with current water will give rise to a large lake of to the shape of the snout and teeth number are also employed to standing water. Fish that live in this type of environment are justify the recognition of these specimens as different species, not adapted to slow flowing waters or lenthic environments but these differences are not constant in each subpopulation. A which are low in oxygen. Furthermore, dozens fish species that similar scenario can be observed for S. zuanoni, with different exist only at the Volta Grande may disappear, because while subpopulations along the rio Iriri (L354). It is not uncommon, the region is a lotic environment, it will not keep sufficient thought, to find further variations on the color pattern of all flow to maintain such species (L. M. Sousa, pers. comm.). aforementioned subpopulations, which hinders the recognition The Xingu Spectracanthicus species are widely distributed of populations as independent species within the genus. along the rio Xingu and have several subpopulations. In this All species studied in the present contribution have case, some of these subpopulations may be greatly reduced or commercial interest as ornamental fish. The ornamental fish even become locally extinct, but the species as a whole will exploration has been a common practice in the rio Xingu and probably not be extinguished, since it will remain in little or rio Tapajós regions for the last 30 years. These activities have non-impacted areas where the water reaming is current. The become economically profitable within some communities same scenario may be expected to occur with S. zuanoni, even of these regions and, in many cases, constitute the entire though its subpopulations are smaller than S. puctatissimus. source of income for several families. According to Rapp The construction of a complex of five dams at rio Tapajós Py-Daniel & Zuanon (2005) and Camargo et al. (2011) this and rio Jamaxim is already scheduled: São Luiz do Tapajós and practice has become most frequent in Altamira since end of Jatobá, on rio Tapajós; and Cachoeira dos Patos, Jamanxim and the1980’s, when gold prospectors that lost their jobs began to Cachoeira do Caí on the rio Jamanxim (Fernandes et al., 2010). collect loricariids fish. The capture of attractive species for the Regarding S. murinus, which has relatively high abundance ornamental fish trade was possible with the adaptation of old in the region of Itaituba, such impacts can directly affect the techniques employed in gold prospect at great depths (up to population due to habitat loss. However, as the species has a 15 meters), such as the use of air compressors, diving masks wide distribution along the rio Tapajós, some subpopulations and lanterns. From this period several loricariid species have will probably not be impacted by habitat loss, being restricted become known worldwide. Presently, the Xingu and Tapajós to regions where the environment remains lotic. Concerning S. drainages together constitute one of the largest sources of immaculatus, although it has little representation in scientific ornamental fish of the family Loricariidae. In the rio Xingu collections, it is abundant at the region and this fact can be the ornamental fishery activities is very intense, especially explaned by the lack of specific capture effort. at Altamira, Senador José Porfírio, Belo Monte, Vitória In rio Tocantins, the Tucuruí Dam construction was do Xingu, and Porto de Moz (Camargo et al., 2011; L. M. completed in 1981 and, as in others dams, the change Sousa, pers. comm.). On the other hand, at rio Tapajós the from a lotic to a lenthic habitat was considered a major ornamental fish activities are concentrated at Santarém and environmental interference, because of the effects to the Itaituba (Sousa & Birindelli, 2009). In the rio Tocantins, the aquatic ecosystem in general. The formation of the lake ornamental fishery is also a common practice concentrated resulted on major changes in the fish fauna of the rio at Marabá region. Tocantins, promoted by disruption of migratory routes and According to Andrews (1990) two of the major ways the disappearance of some species (Mérona et al., 2010). in which the market for ornamental fish can affect natural Although Spectracanthicus tocantinensis is mentioned in populations are the introduction of exotic species and the several web sites of ornamental fish asSpectracanthicus sp. 24 Revision of Spectracanthicus

L086, LDA014, their representation in scientific collections improved by the valuable suggestions and comments of is relatively low and the last samples were collected in 1984. André Netto-Ferreira, Fabio Gaiger, José Birindelli and This decrease of sampling at Tucuruí and Cametá regions two anonymous reviewers. CCC was funded by FAPESP may be related to deficient collection effort for ornamental (proc. 2007/07770-5). fisheries. Furthermore, the species seems to occur in abundance at Marabá region, where it is commercialized Literature Cited for several regions in the country (H. Anatole, pers. comm.) Given the current scenario of investment in hydropower Armbruster, J. W. 2004. Phylogenetic relationships of the suckmouth armoured catfishes (Loricariidae) with emphasis on energy in the Brazilian Amazon, it can be stated that fish the Hypostominae and the Ancistrinae. Zoological Journal of species that are directly dependent on rapid waters with high Linnean Society, 141: 1-80. oxygen content are in threat. Obviously, species with a wide Arratia, G. & L. Huaquin. 1995. Morphology of the lateral line system distribution along certains Amazonian rivers or in several and the skin of diplomystid and certain primitive loricarioid drainages will be safe. On the other hand, species that are catfishes and systematic and ecological considerations. Bonner Zoologische Monographien, 36: 5-109. naturally rare, usually restricted to specific habitats and Andrews, C. 1990. The ornamental fish trade and fish conservation. localities, may suffer a drastic population decline and even Journal of Fish Biology, 37: 53-59. become extinct. Burgess, W. E. & L. Finley. 1996. An atlas of freshwater and marine catfishes: Update. Tropical Fish Hobbyist, 1996: 163-174. Key to the species of Spectracanthicus Camargo, M., J. Carvalho Júnior & R. A. Estupiñan. 2001. Peixes Comerciais da Ecorregião Aquática Xingu-Tapajós. Pp. 175-192. In: Castilhos, Z. C. & P. A. Buckup (Eds.). Ecorregião Aquática 1a. Triangle-shaped opercle lacking large eversible odontodes; Xingu-Tapajós. Rio de Janeiro, CETEM. three anal-fin branched rays………………….S. murinus Camargo, M., H. Gimênes Junior & L. H. Rapp Py-Daniel. 2012. 1b. Bar-shaped opercle with large eversible odontodes; four Acaris ornamentais do Médio Rio Xingu. Belém, FAPESPA/ anal-fin branched rays …...... 2 FUNCEFET. Chamon, C. C. 2012. Revisão taxonômica e relações filogenéticas 2a. Ground color of body dark gray without dots or spots ...... do grupo Acanthicus (Siluriformes, Loricariidae). Unpublished ...... S. immaculatus Ph.D. Dissertation, Universidade de São Paulo, São Paulo, 270p. 2b. Body dark brown to black with small yellowish dots or Eigenmann, C. H. & R. S. Eigenmann. 1889. Preliminary notes on spots, or body dark gray without dots or spots ...... South American Nematognathi. II. Proceedings of the California ...... 3 Academy of Sciences, 2: 28-56. Eschmeyer, W. N. (Ed.). 1998. Catalog of fishes. Special publication, 3a. Body with large (similar to slightly larger eye diameter) California Academy of Sciences, San Francisco. white spots ...... …S. zuanoni Fernandes, F. R. C., G. R. Silva, M. H. M. R. Lima, N. S. Teixeira & 3b. Body with pale/yellowish, small- to medium-sized (clearly R. B. R. Villa Verde. 2001. A Ecorregião Aquática Xingu-Tapajós smaller than eye diameter) dots ...... 4 na Amazônia: Abordagem Demográfica, Econômica e Social. Pp. 4a. Dorsal profile of snout slightly pointed, few premaxillary 59-102. In: Castilhos, Z. C. & P. A. Buckup (Eds.). Ecorregião Aquática Xingu-Tapajós. Rio de Janeiro, CETEM. and dentary teeth (5-8 and 8-16 respectively) ...... Ferraris, C. J. Jr. 2007. Checklist of catfishes, recent and fossil ...... S. tocantinensis (Osteichthyes: Siluriformes), and catalogue of siluriform primary 4b. Dorsal profile of snout rounded, numerous premaxillary types. Zootaxa, 1418: 1-628. and dentary teeth (12-17 and 21-30 respectively) ...... Fisch-Müller, S. 2003. Subfamily Ancistrinae (Armored catfishes)...... S. punctatissimus Pp. 373-400. In: Reis, R.E., S.O. Kullander & C. J. Ferraris Jr. (Eds.). Check list of the Freshwater Fishes of South and Central America. Porto Alegre, Edipucrs, 729p. Acknowledgments Higuchi, H. 1996. An updated list of ichthyological collecting stations of the Thayer Expedition to Brazil (1865-1866). Available Authors wish to thank Jonathan Armbruster from: http://www.mcz.harvard.edu/Departments/Fish/thayer. (AUM), Jansen Zuanon (INPA), Sonia Fisch-Müller htm. (May 2013). Fowler, H. W. 1914. Fishes from the Rupununi River, British Guiana. (MHNG), Marcelo Britto and Paulo Buckup (MNRJ), Proceedings of The Academy of Natural Science of Philadelphia, Helmut Wellendorf (NMW), Michel Gianetti and Osvaldo 66: 229-284. Oyakawa (MZUSP) and Hielke Praagman (ZMA) for Isbrücker, I. J. H. 1980. Classification and catalog of the mailed the hospitality during visits to the fish collections under Loricariidae (Pisces, Siluriformes). Verslagen en Techniche their care, for loan of specimens provide collection data Gegevens, Instituut voor Taxonomische Zoölogie, Universiteit van Amsterdam, 22: 1-181. information. Thanks are also due to José Birindelli (UEL) Isbrücker, I. J. H. & H. Nijssen. 1991. Hypancistrus zebra, a new and Leandro Sousa (UFPA) for permission to use their genus and species of uniquely pigmented ancistrine loricariid fish images of live specimens, and Eduardo Baena for the from the rio Xingu, Brazil (Pisces: Siluriformes: Loricariidae). osteological structures images. The manuscript was also Ichthyological Exploration of Freshwaters, 1: 345-350. C.C. Chamon & L.H.R. Py-Daniel 25

Kner, R. 1853. Die Hypostomiden. Zweite Hauptgruppe der familie Schaefer, S. A. 1987. Osteology of der Panzerfische. (Loricata vel Goniodontes). Denkschriften der (Linnaeus), with a phylogenetic analysis of the loricariid Kaiserlichen Akademie der Wissenschaften, 7: 251-286. subfamilies (Pisces, Siluroidei). Contributions in Sciences, Lujan, N. K., M. Arce & J. W. Armbruster. 2009. A new black Natural History Museum of Los Angeles County, 394: 1-31. Baryancistrus with blue sheen from the upper Orinoco Schaefer, S. A. 1986. Historical Biology of the loricariid catfishes: (Siluriformes: Loricariidae). Copeia, 2009: 50-56. phylogenetics and functional morphology. Unpublished Ph.D. Lundberg, J. G. & J. N. Baskin. 1969. The caudal skeleton of the Dissertation, The University of Chicago, Chicago, 290p. catfishes, order Siluriformes. American Museum Novitates, Schraml, E. & F. Schafer. 2004. Aqualog: Loricariidae All L-Numbers. 2398: 1-49. Rodgau, Hollywood Import & Export, Inc., 2nd Edition. Mérona, B. Juras, A. A., Santos, G. M. & I. H. A. Cintra. 2010. Sousa, L. M. & J. L. O. Birindelli. 2009. Acaris do Tapajós. Habitat, Os peixes e a pesca no rio Tocantins: vinte anos depois da 84: 8-19. UHE Tucuruí. Brasília, Centrais Elétricas do Norte do Brasil, Steindachner, F. 1881. Beitrage zur Kenntniss der Flussfische Eletronorte. Su damerika’s. II. Denkschriften der Mathematisch- Rapp Py-Daniel, L. H. 1989. Redescription of Parancistrus Naturwissenschaftlichen Classe der Kaiserlichen Akademie der aurantiacus (Castelnau, 1855) and preliminary establishment of Wissenschaften in Wien, 43: 103-146 two new genera: Baryancistrus and Oligancistrus (Siluriformes, Taylor, W. R. & G. C. Van Dyke. 1985. Revised procedures for Loricariidae). Cybium, 13: 235-246. staining and clearing small fishes and other vertebrates for bone Rapp Py-Daniel, L. H. & J. A. Zuanon. 2005. Description of a new cartilage. Cybium, 9: 107-119. species of Parancistrus (Siluriformes: Loricariidae) from the rio Werner, A., W. Lechner & J. Schmidt. 2005. Miniatlas - L-Welse - L1 Xingu, Brazil. Neotropical Ichthyology, 3: 571-577. bis L400. Ruhmannsfelden, Bede-Verlag. Reis, R. E., E. H. L. Pereira & J. Armbruster. 2006. Delturinae, a Zuanon, J. A. 1999. História natural da ictiofauna de corredeiras do rio new loricariid catfish subfamily (Teleostei: Siluriformes), with Xingu, na região de Altamira, Pará. Unpublished Ph.D. Dissertation, revisions of Delturus and Hemipsilichthys. Zoological Journal Universidade Estadual de Campinas, Campinas, 199p. of Linnean Society 147: 277-299. Reis, R. E., S. O. Kullander & C. J. Ferraris. 2003. Family Submitted August 4, 2012 Loricariidae. Pp. 318. In: Reis, R. E., S. O. Kullander & C. J. Accepted November 19, 2013 by Francisco Langeani Ferraris. (Eds.). Checklist of the freshwater fishes of South and Central America. Porto Alegre, Edipucrs. Published March 31, 2014